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Pattern Formations with a Zooplankton: Experiment and Theory. Anke Ordemann Frank Moss. Support:. US Office of Naval Research Alexander von Humboldt Foundation. Motivation: Vortex-swarming of ‘self-propelled’ animals. Blue Planet: Seas of Life, Discovery Channel 2001. Outline.
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Pattern Formations with a Zooplankton: Experiment and Theory Anke Ordemann Frank Moss Support: US Office of Naval Research Alexander von Humboldt Foundation
Motivation:Vortex-swarming of ‘self-propelled’ animals Blue Planet: Seas of Life, Discovery Channel 2001
Outline • The Active Brownian Particle (ABP) Theory – Erdmann, Ebeling, Schweitzer – Humboldt University Berlin • Single (few) Particle Motions & Bifurcations • About Daphnia – the experiment • Random Walk Theory - Measures and Results • Symmetry Breaking – Vortex motion • Simulation using RWT • Summary
Active Brownian Particle (ABP)with internal energy depot [Schweitzer et al., PRL 80, 5044 (1998)] Energy depot: space-dependent take-up q(r), internal dissipation ce(t), conversion of internal energy into kinetic energy d2e(t)v2 (For uniform distr: q(r) = q0) A velocity dependent dissipation
Hopf bifurcation: Fixed point limit cycle pair Bifurcation parameter:
Daphnia make vortices Side view of vortex-swarming zooplankton Daphnia in our lab. Picture by D.F. Russell. Schlieren technique movie of Daphnia motion by Ai Nihongi and Rudi Strickler.
About Daphnia Daphnia magna, common “water flea”. • They are: • Attracted to visible light (VIS) • Repelled by ultra violet (not used here) • Blind to infra red (IR) Record their motions using IR and manipulate motions using VIS
Apparatus • Side and bottom views recorded simultaneously • VIS light shaft is the cue • Illumination and recording in IR
Example RecordingsAbout 40 animals, approximately equal numbers circling in opposite directions Bottom view of a few Daphnia moving around light shaft (radius r =5mm). Movie is speeded up 4 times. Five successive positions of Daphnia taken in intervals of 0.3s (black/white inverted).
Observation of a single Daphnia Note reversal X(t) Y(t) Bottom view of individual Daphnia moving around light shaft. Movie is speeded up 3 times. Track of one Daphnia individually circling horizontally around light shaft (146s). Single Daphnia circle individually in both directions around light shaft, frequently changing direction
How to characterize observed circular motion?- M=624 moves from 4 different animals - Average speed vavg = 5.71 ± 1.35 mm/s MC= Number of successive moves before changing direction (CCW vvs CW). MC = 11.8
Random Walk Theory - but needMotion of single Daphnia in the dark. 1599 moves from 8 different animals Example track, 200 hops X Distribution of turning angles, = directional change after each hop Y P() symmetric about 0. || 35 (Similar to oceanic copepods)
Random Walk Theory *Short range temporal correlation *Attraction to light • Daphnia in darkness: At the end of each step, the direction of next time step at ti+1 is randomly chosen from observed distribution of turning angle (DTA) • Daphnia in light field: At the end of each time step, the direction is again chosen from the DTA, but an additional kick of strength r*L/(L-1) towards the light (parabolic potential) is added and the final heading is rescaled to unit length. [0 L< 1]
Results of the Simulation for various L: P(θ) Heading angle, P(r) Radius, r
Results (cont) MC L P(MC) L = 0.4 MC
Many Particle Generalizations - ABP (Humboldt) Particle-Particle attractive interaction Mean field potential (ABP) and probability densities of the limit cycle motions. Symmetric pair of limit cycles. Interacting Active Brownian Particles[Schweitzer et al., PRE 64, 021110 (2001); Ebeling and Schweitzer, Theory Biosci. 120, 20 (2001); Erdmann et al., PRE 65, 061106 (2002)]
Many Particle Generalizations – Levine Short range repulsion and long range attraction added to the model of Self-Propelled Interacting Particles [Levine et al., PRE 63, 017101 (2001)] Self propelling force, (no alignment) Self propelling force, (with alignment – range lc)
Transition to Vortex Motion by a Daphnia SwarmModels:Symmetry of limit cycles must be broken.Velocity alignment; Avoidance Daphnia experiment:hydrodynamic coupling likely(for birds and fish the alignment is visual with neighbors) CCW Motion and spiral arms
Another example CW Motion (after a time delay)
RWT Simulation of the transition: Interacting RW with DTA • Simple model for (vortex-) swarming Daphnia • Indirect inter-agent interactions via water drag are incorporated by adding ‘alignment’ or ‘water drag’ kick proportional to: A local order parameter: Ntot = 30 O Time, t Global order parameter: O = (NCW NCCW )/Ntot
Conclusions Theory predicts four motions: • Noisy fixed point • Symmetric pair of limit cycles • Swarming • Transition to vortex All four motions can be observed in experiments with Daphnia
Discussion What are the minimum ingredients that lead to these motions? A. Few animal (low density) experiments: • Self-propelled particles (finite non-zero velocity) • Assigned preference to move in ‘forward’ direction • Confinement, arising from Attraction (either as mean field potential from interagent interactions or as external attractive potential) B. Large animal density experiments: • In addition to the above – a symmetry breaking (velocity alignment) mechanism.